Viverravidae

Viverravidae is an extinct family within the clade Carnivoramorpha. They are related to carnivorans, and lived from the early Palaeocene to the Eocene.

Viverravidae Temporal range: 66.043–33.9 Ma PreꞒ Ꞓ O S D C P T J K Pg N early Paleocene - late Eocene
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Clade:	Carnivoramorpha
Superfamily:	†Viverravoidea Wortman & Matthew, 1899
Family:	†Viverravidae Wortman & Matthew 1899[1]
Genera
[see classification]
Synonyms
Didymictidae[2]

In viverravids, the number of molars is reduced to two and the skull is elongated. Viverravidae is a monophyletic family, a plesion-group. They are not thought to be ancestral to any extant carnivorans.[3]

The viverravids were thought to be the earliest carnivorans: they first appeared in the Paleocene of North America about 60 million years ago. One author proposed that they should be placed outside the order Carnivora based on cranial morphology.[4]

Wang and Tedford propose that they arose in North America 65-60 million years ago, spread to Asia then later to Europe, and were the first carnivorans and possessed the first true pair of carnassial teeth.[5]^:p8

Classification and phylogeny

Classification

Taxonomy retrieved from the Paleobiology Database[6]

• Superfamily: †Viverravoidea (Wortman & Matthew, 1899)
  • Family: †Viverravidae (Wortman & Matthew, 1899) [synonym: Didymictidae (Flynn & Galiano, 1982)]
    • (unranked): †Ictidopappus/Pappictidops clade
      • Genus: †Orientictis (Huang & Zheng, 2005)
        • †Orientictis spanios (Huang & Zheng, 2005)
      • Genus: †Pappictidops (Qiu & Li, 1977)
        • †Pappictidops acies (Wang, 1978)
        • †Pappictidops obtusus (Wang, 1978)
        • †Pappictidops orientalis (Qiu & Li, 1977)
      • Genus: †Variviverra (Tong & Wang, 2006)
        • †Variviverra vegetatus (Tong & Wang, 2006)
      • Subfamily: †Ictidopappinae (Van Valen, 1969)
        • Genus: †Ictidopappus (Simpson, 1935)
          • †Ictidopappus mustelinus (Simpson, 1935)
    • Subfamily: †Didymictinae (Flynn & Galiano, 1982)
      • Genus: †Bryanictis (MacIntyre, 1966)
        • †Bryanictis microlestes (Simpson, 1935)
        • †Bryanictis paulus (Meehan & Wilson, 2002)
      • Genus: †Didymictis (Cope, 1875)[7]
        • †Didymictis altidens (Cope, 1880)
        • †Didymictis dellensis (Dorr, 1952)
        • †Didymictis leptomylus (Cope, 1880)
        • †Didymictis protenus (Cope, 1874) [synonym: Didymictis curtidens (Cope, 1882)]
        • †Didymictis proteus (Polly, 1997)
        • †Didymictis vancleveae (Robinson, 1966)
      • Genus: †Intyrictis (Gingerich & Winkler, 1985)
        • †Intyrictis vanvaleni (MacIntyre, 1966)
      • Genus: †Protictis (Matthew, 1937)
        • †Protictis agastor (Gingerich & Winkler, 1985)
        • †Protictis haydenianus (Cope, 1882) [synonyms: Didymictis primus (Cope, 1884), Prolimnocyon macfaddeni (Rigby, 1980)]
        • †Protictis minor (Meehan & Wilson, 2002)
        • †Protictis paralus (Holtzman, 1978)
        • †Protictis simpsoni (Meehan & Wilson, 2002)
      • Genus: †Protictoides (Flynn & Galiano, 1982)
        • †Protictoides aprophatos (Flynn & Galiano, 1982)
      • Genus: †Raphictis (Gingerich & Winkler, 1985)
        • †Raphictis gausion (Gingerich & Winkler, 1985)
        • †Raphictis iota (Scott, 2008)
        • †Raphictis machaera (Rankin, 2009)
        • †Raphictis nanoptexis (Rankin, 2009)
    • Subfamily: †Viverravinae (Wortman & Matthew, 1899)
      • Genus: †Pristinictis (Fox & Youzwyshyn, 1994)
        • †Pristinictis connata (Fox & Youzwyshyn, 1994)
      • Genus: †Simpsonictis (MacIntyre, 1962)
        • †Simpsonictis jaynanneae (Rigby, 1980)
        • †Simpsonictis pegus (Gingerich & Winkler, 1985)
        • †Simpsonictis tenuis (Simpson, 1935)
      • Genus: †Viverravus (Marsh, 1872) [synonym: Ziphacodon (Marsh, 1872)]
        • †Viverravus acutus (Matthew & Granger, 1915)
        • †Viverravus gracilis (Marsh, 1872) [synonyms: Didymictis dawkinsianus (Cope, 1881), Harpalodon vulpinus (Marsh, 1872), Triacodon fallax (Marsh, 1871), Ziphacodon rugatus (Marsh, 1872)]
        • †Viverravus lawsoni (Hooker, 2010)
        • †Viverravus laytoni (Gingerich & Winkler, 1985) [synonym: Viverravus bowni (Gingerich, 1987)]
        • †Viverravus lutosus (Gazin, 1952)
        • †Viverravus minutus (Wortman, 1901)
        • †Viverravus politus (Matthew & Granger, 1915) [synonym: Protictis schaffi (Gingerich & Winkler, 1985)]
        • †Viverravus rosei (Polly, 1997)
        • †Viverravus sicarius (Matthew, 1909)
      • Genus: †Viverriscus (Beard & Dawson, 2009)
        • †Viverriscus omnivorus (Beard & Dawson, 2009)

Phylogeny

The phylogenetic relationships of Viverravidae are shown in the following cladogram:[8][9][10][11][12][13]

Carnivoramorpha

Carnivoraformes    ?  †carnivoramorph sp. (UALVP 31176)    ?  †carnivoramorph sp. (WW-84: USNM 538395)      ?  †Ravenictis    ?  †carnivoramorph sp. (UALVP 50993 & UALVP 50994)    †Viverravoidea   †Viverravidae         †Variviverra  †Variviverra vegetatus  ?   †Orientictis  †Orientictis spanios      †Pappictidops    †Pappictidops orientalis       †Pappictidops acies     †Pappictidops obtusus          †Ictidopappinae   †Ictidopappus    †Ictidopappus mustelinus     †Ictidopappus sp. (AMNH 58409)       †Ictidopappus/Pappictidops група      †Viverravinae     †Simpsonictis      †Simpsonictis pegus     †Simpsonictis tenuis       †Simpsonictis sp.     †Simpsonictis jaynanneae      †Viverriscus  †Viverriscus omnivorus      †Pristinictis  †Pristinictis connata  ?   †Viverravus      †Viverravus gracilis     †Viverravus lutosus     †Viverravus minutus     †Viverravus sicarius       †Viverravus politus     †Viverravus lawsoni       †Viverravus laytoni       †Viverravus acutus     †Viverravus rosei            ?  †viverravid sp. (CM 71188 & CM 71189)    †Didymictinae     †Raphictis    †Raphictis gausion     †Raphictis iota     †Raphictis machaera     †Raphictis nanoptexis     †Raphictis sp. (UALVP 51558)     †Raphictis sp. (UALVP 8861)       †viverravid sp. (UALVP 51560)      †Didymictis      †Didymictis leptomylus     †Didymictis protenus       †Didymictis vancleveae     †Didymictis altidens     †Didymictis sp. (YPM VPPU 024902)    †Didymictis proteus    †Didymictis proteus (sensu stricto)    ?  †Didymictis dellensis    sensu lato     †Protictis     †Protictis      †Protictis paralus     †Protictis agastor       †Protictis simpsoni     †Protictis minor     †Protictis haydenianus    sensu stricto   †Protictoides  †Protictoides aprophatos      †Bryanictis   †Intyrictis  †Intyrictis vanvaleni    †Bryanictis    †Bryanictis microlestes     †Bryanictis paulus     †Bryanictis sp. (Dawson Creek)    sensu stricto   sensu lato   sensu lato

References

1. Wortman, J. L.; Matthew, W. D. (1899). "The ancestry of certain members of the Canidae, Viverridae, and Procyonidae". Bulletin of the American Museum of Natural History. 12: 109–138. hdl:2246/1535. OCLC 46687698.CS1 maint: ref=harv (link)
2. "†family Viverravidae (Wortman & Matthew, 1899) (placental)". Fossilworks. Retrieved 29 June 2019 from the Paleobiology Database.
3. Wesley-Hunt, G. D.; Flynn, J. J. (2005). "Phylogeny of the Carnivora: basal relationships among the carnivoramorphans, and assessment of the position of 'Miacoidea' relative to Carnivora". Journal of Systematic Palaeontology. 3: 1–28. doi:10.1017/S1477201904001518.
4. Polly, David, Gina D. Wesley-Hunt, Ronald E. Heinrich, Graham Davis and Peter Houde (2006). "Earliest known carnivoran auditory bulla and support for a recent origin of crown-clade carnivora (Eutheria, Mammalia)" (PDF). Palaeontology. 49 (5): 1019–1027. doi:10.1111/j.1475-4983.2006.00586.x.CS1 maint: multiple names: authors list (link)
5. Wang, Xiaoming; Tedford, Richard H.; Dogs: Their Fossil Relatives and Evolutionary History. New York: Columbia University Press, 2008.
6. Paleobiology Database. Retrieved with Fossilworks (March 3, 2017)
7. Zack, Shawn P. (2012). "Deciduous dentition of Didymictis (Carnivoramorpha: Viverravidae): implications for the first appearance of "Creodonta"". Journal of Mammalogy. 93 (3): 808–817. doi:10.1644/11-MAMM-A-245.1. ISSN 0022-2372.
8. J. J. Flynn and H. Galiano. (1982.) "Phylogeny of Early Tertiary Carnivora, With a Description of a New Species of Protictis From the Middle Eocene of Northwestern Wyoming" American Museum Novitates 2725:1-64
9. P. D. Gingerich and D. A. Winkler. (1985.) "Systematics of Paleocene Viverravidae (Mammalia, Carnivora) in the Bighorn Basin and Clark's Fork Basin, Wyoming." Contributions from the Museum of Paleontology, University of Michigan 27(4):87-128
10. P. D. Polly. (1997.) "Ancestry and Species Definition in Paleontology: A Stratocladistic Analysis of Paleocene-Eocene Viverravidae (Mammalia, Carnivora) from Wyoming." Contributions from the Museum of Paleontology, University of Michigan 30(1):1-53
11. Malcolm C. McKenna, Susan K. Bell: Classification of Mammals: Above the Species Level in Columbia University Press, New York (1997), 631 Seiten. Viverravidae
12. Solé, Floréal; Smith, Thierry; De Bast, Eric; Codrea, Vlad; Gheerbrant, Emmanuel (2016). "New carnivoraforms from the latest Paleocene of Europe and their bearing on the origin and radiation of Carnivoraformes (Carnivoramorpha, Mammalia)". Journal of Vertebrate Paleontology. 36 (2): e1082480. doi:10.1080/02724634.2016.1082480. ISSN 0272-4634.
13. S. Faurby, L. Werdelin, A. Antonelli (2019.) "Dispersal ability predicts evolutionary success among mammalian carnivores" Department of Biological and Environmental Sciences, University of Gothenburg, Box 461, SE